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Plants mate. In a manner far more elegant than our own mammalian shenanigans, and far more important for the ongoing survival of the Earth’s ecosystems, plants are out there constantly having sex. And it is a deeply interesting thing involving insects, and wind, and stigmas, and stamens, and enough puzzling evolutionary biology to fill journal articles and occupy a small handful of academic careers.

My latest paper is about plant mating, and rather than wade through meadows of blooms to collect the data for this effort, I remained desk-bound in Milwaukee, wading through 30 years of plant mating literature with the intention of assembling a dataset to help us see plant mating in a new light.

A mint-bush (Prostanthera lasianthos), caught in the act of mating.

As you may know, the flowers of many plant species are hermaphrodite, i.e. they bear both male and female sexual organs. This means plants can have sex with themselves. It’s called self-pollination, or “selfing”, and it can happen in two ways. First, pollen can move from the male parts of a flower to the female parts of the same flower. Second, on a single plant, pollen can move from the male parts of one flower to the female parts of a different flower.

“But isn’t this inbreeding??” I hear you ask, recoiling in horror at the moral and population genetic gutter down which I have so suddenly led you. Yes! Seed fertilized in a selfed flower is about as inbred as it gets, even more so than sibling mating. But you see, inbreeding is not always a bad thing—in fact inbreeding can actually be beneficial in some circumstances—and here’s where things get messy.

Genes are selfish. As such, they will do whatever it takes to get themselves propagated into another body. Consider then the selfish genes of a mother plant, which want to get into the next generation, and into as many individuals as possible. If a mother plant mates with a genetically different plant (an “outcross” mating event), its seeds each inherit approximately 50% of the mother’s genetic variation. This is because they combine with (and are diluted by) the genes of the pollen donor during pollination and sexual reproduction. On the other hand, if a mother self-fertilizes its own seeds, it transmits closer to 100% of its genetic legacy into each offspring. In the eyes of natural selection, this is a huge difference, and offers a great advantage to selfing over outcrossed reproduction.

So why aren’t all plants selfing all the time? This is because the huge advantage in transmitting genes via selfing is offset by the drawbacks of inbreeding. You can’t often get away with inbreeding without cost, and the cost is that inbred offspring are commonly less fit than their “outbred” siblings. Called “inbreeding depression”, this occurs because breeding from related individuals vastly raises the probability of combining rare genetic variants that harm the individual they are in. In the long run, inbreeding also reduces genetic variation that is essential for adapting to changing environments.

Evolutionary biologists have been pondering this tension for decades, and it led to the hypothesis that plants should be driven by natural selection into two polar opposite strategies: selfing plants should be favoured by natural selection when inbreeding depression is weak, and outcrossing plants should be favoured by natural selection when inbreeding depression is strong. Anything in the middle should not be adaptive, and should therefore be rare in nature. Sounds sensible, except numerous studies have now measured selfing/outcrossing across hundreds of plant species, and there is a curious and difficult-to-explain preponderance of plants that are neither exclusively selfing or outcrossing. These plants are having it both ways, seemingly hedging between strategies, and we call them the “mixed maters”.

Now for the problem my study addresses… Over the last 30 years, loads of studies have been measuring the selfing/outcrossing rate of plants in the wild. There have also been important studies which collect all these outcrossing estimates into big global datasets and generate observations like the one above: that mixed-mating is inexplicably common. For 30 years however, much of this discussion on global patterns was centred around average outcrossing per species, and using this to classify a species as a “selfer”, “outcrosser”, or “mixed mater”. We know evolution doesn’t work on a “species level” though. What is more appropriate is what is happening in a population context. And perhaps by averaging away all the variation within each species, my co-authors and I thought we might be missing an important perspective on the data.

What my co-authors and I did then was to collect all the outcrossing estimates from published papers containing three or more population outcrossing estimates. Going back 30 years, we collected data for 105 species and measured the variation in outcrossing within a species, among populations.

What we found was huge variation! There was commonly so much variation in mating within a species, that species averages felt inadequate for expressing where mating system variation lies. The data also showed that variation was difficult to predict. For example, there was an old hypothesis floating about that wind-pollinated plants were less variable in their outcrossing/selfing rate than animal pollinated species. The reason being that animals were thought to fluctuate more in their abundance and service between sites and seasons, while wind was a more consistent, reliable force for pollination. For the first time we were able to test this, and our analysis did not support the hypothesis. We also tested whether mating variation was evolving in a predictable fashion, but found that the relationship of plant species had no bearing on the variation we found in those species’ mating patterns.

Arabis alpina: revealed in our study as a species with one of the most variable mating systems measured. By Hedwig Storch CC BY-SA 3.0, from Wikimedia Commons.

Ultimately, our study can’t solve the mystery of why we have so many mixed-mating plants. What it does do though is present a new way to look at the known range of variation in plant mating. There are other analyses waiting to be done with the data collected here, and we have highlighted where more studies are needed to answer interesting questions. For example, do different kinds of animal pollinators result in more or less variable outcrossing? Or do populations on the fringe of a species’ range experience more or less variable outcrossing?

As the body of knowledge on plant mating systems continues to grow, this dataset will grow too, and this population-level perspective on plant mating will hopefully provide the basis for the next insights into evolution’s influence on plant mating.

Earlier this year, I decided A. stomarrhena looked like a perfect candidate for my new study on pollinators and gene flow. What I needed was a bird-pollinated species of plant, closely related to an insect-pollinated species. This one seemed to match all the criteria I needed, except there was no evidence that it was bird-pollinated. But with those long, tapered corolla tubes, and that pink-red coloration, I believed that birds absolutely had to be the pollinator.

The danger was, that while birds might be visitors, the plant could be somewhat “generalized”, and also use insects. This is pretty common, especially in places like Australia where European Honeybees (Apis mellifera) have invaded ecosystems that evolved in their absence, and honeybees will visit absolutely everything whether the plants are adapted to bees or not.

By deploying a new camera-trapping method that I am developing to record insect visitation, I was able to gather several days of pollinator observations, despite some very bad weather. After initially being baffled as to what honeyeater might visit such a low ground-hugging shrub, I got my answer after day one, when I captured video of my new favourite bird: the Tawny-crowned Honeyeater (Gliciphila melanops) feeding on the flowers. Furthermore, the recordings of honeybee fly-bys are sufficient to rule them out as pollinators.

This little result is a win on two fronts: a successful trial of new pollinator-monitoring cameras, and vindication of predicting pollinators from flower morphology.

Rare plants nurseries are like second hand bookshops. It’s always so tempting to browse on the off chance you find that little treasure. I recently visited a charming rare plants nursery in Mt Macedon (boutique-y town outside Melbourne, Australia) where I discovered these for sale:

Hello old friend! (Hesperantha coccinea)

The last time I saw this elegant iris, it was flowering on stream banks 10,000 km away in the Drakensberg Mountain range in South Africa. There in its natural habitat, it is pollinated in some areas by a very special butterfly: the Mountain Pride (Aeropetes tulbhagia). In other places, it is pollinated by the amazing long-tongue fly (Prosoeca ganglbaueri). The two forms are a wonderful example of “pollination ecotypes”, where different populations are undergoing adaptation to their unique pollinators. The fly-serviced ones are a pink hue with narrow petals, while the butterfly-pollinated ones are much redder with broader petals.

Hesperantha coccinea at home in South Africa with its pollinator (Prosoeca ganglbaueri).

Fast forward two weeks, and I’m home walking the dog in my quite unremarkable Melbourne suburb, when who should I see?

Hello old friend! (Diascia sp.)

It’s winter here, with very little in flower, but these brilliant little pink blooms volunteering themselves from underneath a fence in suburban Melbourne really made my day. The last time I saw a Diascia, it was growing amongst the boulders on creek beds and on cliffs in the Drakensberg Mountains. These are Diascia, or “twinspur” and its this common name that alludes to their fascinating pollination story.

Hug-pollination by oil-collecting bee (Rediviva sp.) in Diascia.

Diascia have two spurs on the back of the flower, which is distinct from the usual arrangement of a single nectar-spur. The difference is that these flowers don’t reward pollinators with sugary secretions, instead they provide oil to specialised oil-collecting bees in the genus Rediviva. The bees use this oil to line their nests and provision their young. In order to collect the nectar, they must reach deep into the twin spurs with their lanky forelimbs, and comb it out. In so doing, they effectively hug the reproductive parts of the Diascia flower and effect pollination.

In Spring, I plan to take some cuttings from this little Diascia. Keeping species with special personal significance is a deeply satisfying part of cultivating plants. A plant can be kept like a souvenir or memento marking a time in one’s life, just like a photo or trinket. But plants have an advantage over these inanimate reminders. Because biological reproduction requires the physical donation of part of the mother’s cells to the daughter cells, my keepsake plant can be viewed as a physical part of the plant that appears in my fond memory. If I could see in four dimensions, I could literally look down the line of cell-divisions all the way back to where the Hesperantha in the nursery physically intersects as the same individual with the Hesperantha I observed flowering in the Autumn sun of the Drakensberg Mountains in South Africa.

Let’s say, however, that your orchid parents do manage to achieve fertilization. Your orchid mother will produce many thousands of tiny dust-like seed, which will be jettisoned into the wind. Unlike most seeds, you have no maternal energy investment to power your germination and first days as a seedling. Instead, you must rely on blind luck to land you within reaching distance of a strand of soil fungus. This fungus is the wet nurse to bring you into the world, invading the seed coat and hooking the young orchid up to a network of fungal strands that pervade the soil. Tapping into this network provides you with the first sips of carbohydrate and nutrient you need in order to build your first green leaf and begin to stand on your own roots. But it is not enough to land near any fungus. Many orchid species require fungal partnership with a specific species of fungus for this to occur at all. Multiplied together, it is a wonder that orchids ever overcome these odds to propagate themselves into the next generation.

The southwest of Western Australia is rightly famous as a global biodiversity hotspot. The area is particularly rich in orchids, and the spider orchids (Caladenia) are some of the most impressive and diverse of the region’s main orchid groups. In 1967, University of Adelaide researcher John Warcup discovered in association with Caladenia a new genus of fungi. Today those fungi are called Serendipita, and although we have known of them for around 60 years, there have been less than a handful of species discovered and described.

The spider orchid Caladenia arenicola was one of those sampled in the study

White spider orchid (Caladenia splendens)

Ubiquitous yet invisible

Although related to mushrooms, Serendipita fungi have not been observed producing the conspicuous spore-bearing fruit bodies we usually use to find and identify them. This makes them largely invisible, and I have therefore never observed them in the wild. Despite that, recent research using DNA sequencing has found them to be absolutely everywhere. Inside all kinds of plants, outside all kinds of plants, and distributed from the equator to Antarctica. It is clear then that there must be a hidden biodiversity of these species siting, waiting to be discovered.

My study took a wide sample of southwest WA spider orchid samples and assayed them for the presence of Serendipita fungi. We then sequenced the DNA of all the fungi we found, and used a new analytical technique for dividing that DNA sequence diversity into units that are probably species. This is currently the only way to sensibly identify Serendipita fungi, as they all look completely alike and do not produce spores in the lab.

We found a total of eight species of Serendipita fungi, including the original species discovered by Warcup back in the 60s. These came from a total of 18 species of orchid. At some sites where we sampled multiple orchid species, we found six species of Serendipita, meaning that the fungi were as diverse as the orchids!

Lying just below the soil horizon, that swollen, yellow stem bit is called the “collar”, and its where all spider orchids keep their fungus.

Untapped agricultural potential?

Although we have chosen to study these Serendipita in association with orchids, their wide host association has got other researchers interested in their role in plant health and application to agriculture. For example, Warcup’s species and one other have been used in experiments (and patent applications) showing inoculation with Serendipita results in profound benefits for the host plant, including:

These proven benefits make Serendipita a potentially powerful tool to enhance plant productivity and stress tolerance in crops. Furthermore, application of Serendipita fungi could be an organic alternative permitting growers to lower the application of unsustainable and ecologically harmful synthetic fertilizers. Our knowledge of plant-Serendipita associations in the wild suggests that these relationships are more prevalent in nutrient poor soils such as those in southwest WA. They are probably one factor that allows our plant diversity to thrive in such weathered, poor soils. This means that species of fungi that have evolved with the nutrient poor soils (like those discovered in this paper) might be untapped tools to enhance agriculture taking place in those very same soils.

(Erratum: This story was edited to replace the figure attributed to Ray and Craven (2016). The first image I used was one showing Arabidopsis capability for mycorrhizal association. Arabidopsis is typically thought to be a non-mycorrhizal plant, which is why this is interesting. The image however showed slower growth in the mycorrhizal treatment. A related Serendipita has been shown to enhance root growth in Arabidopsishowever. I have now updated the post with a more appropriate image of root growth gains in wheat. Thanks to Pawel Waryszak (@PWaryszak) for pointing this out.)

I recently began a brand new project with the University of Melbourne. The beginning of a new project is filled with equal parts excitement and trepidation—excitement at the novelty, the blank canvas, the potential, and trepidation at not wanting to put a foot wrong in critical early decisions that will affect the outcome of a career-defining opportunity.

Here the photos from a first foray into East Gippsland, surveying sites for bird-pollinated Prostanthera walteri.

I recently wrote an article for Wildlife Australia about Australian sexually deceptive orchids, their evolutionary biology, and historical and current research about them. You can download and read the article here: PDF. Thanks to Carol Booth for her collaboration and editorial guidance.

The latest of Australia’s sexually deceptive orchids that I have seen (below) are Caleana major, the Flying Duck orchid (left), and a spider orchid Caladenia clavigera (right). Both were photographed last week in Brisbane Ranges NP, Victoria.

Flowering this year is one of the best seasons of recent times both east and west of the country. So if you’re in Australia, don’t miss the chance to get out bush and enjoy it.

Rather than luring its pollinator with the promise of food this flower uses an equally, if not more, powerful motivator: sex.

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In shades of dusky green and claret red, the bird orchid’s subdued palette hints at its alternative lifestyle. The usual strategy for flowers attempting to catch the compound eye of a passing insect is to advertise proudly. Petals are used as panels for saturated colour, assembled en masse into conspicuous aggregate displays exuding exotic scents. In this way, nectar-filled flowers loudly broadcast the promise of their reward to entice would be pollinators into servicing them.

A deviant among flowering plants, the bird orchid eschews these typical hallmarks of floral advertisement. Crouched modestly on the forest floors of eastern Australia, its stature belies its status as one of the supreme specialists amongst the world’s flowering plants. Like those other showy flowers, the bird orchid needs the service of a pollinator from time to time, however unlike most other flowers, it attracts its pollinator without the payment of any reward. The orchid flower in fact completely lacks nectar.

Rather than luring its pollinator with the promise of food this flower uses an equally, if not more, powerful motivator: sex. Undetectable to human senses, the orchid’s advertisement is a precise chemical mimicry of a female wasp’s sex pheromone. This is targeted marketing at its finest, as the use of a signature sex pheromone ensures that the orchid attracts only males of a specific species of wasp.

Skimming by on wide zig-zagging flights, the wasps are interminably attracted when the ruse takes hold. They alight onto the flower with fervor, probing and hunting for the mate that their senses scream must be there. Bucking back into the column of the flower (the reproductive parts of an orchid flower are fused in this special structure), they make contact with the anthers and a large packet of pollen is deposited on them. The wasp disengages eventually and leaves, but soon, elsewhere, he will catch on the breeze the smell of a mate, and if fooled again, fulfill his role as duped courier for an orchid’s reproductive ends.

Called “sexual deception”, this mode of pollination was noticed by Darwin and his contemporaries in an age in which Europe’s natural sciences were in full bloom. It was a naturalist in Blackburn, Victoria however, who was first to discover the phenomenon outside Europe. In 1927, Edith Coleman had turned her great capacity for observation of the natural world to a peculiar native orchid. Resembling more flesh than flower, Cryptostylis, known also as “tongue-orchids” had caught her attention for its magnetic allure to a specific kind of wasp. Through her observations, Coleman was able to discern that male wasps were being attracted to the flower in order to copulate with it. An experiment through a window showed scent to be the primary attractant, and Coleman even observed the ejaculate remaining after having been visited by clearly convinced wasps. She wrote up her notes in a series of papers for the Victorian Naturalist and Transactions of the Royal Society for Entomology, which made quite a splash with the best of botany at the time.

We now know this was the tip of the iceberg. Australia is not only home to tongue orchids, but hosts a diverse array of other sexually deceptive orchids including the spider orchids, elbow orchids, hammer orchids, dragon orchids, greenhoods, duck orchids, hare orchids, beard orchids, bird orchids, and the list goes on. Harbouring over 50% of the world’s known examples of sexually deceptive pollination, Australia is certainly the world’s hotspot for this unusual phenomenon. Remarkably, we have several hundred species that employ this unique brand of pollinator attraction, and what is more remarkable, the evidence points to at least six different independent evolutionary occurrences in the Australian orchid family tree. To our eyes, sexual deception seems like a freaky, unlikely strategy and its repeated independent incidence through Australia’s evolutionary history is therefore a startling paradox.

Although the reliance on a single species of pollinator for pollination seems precarious, studies have demonstrated that sexual deception comes with the advantage of promoting healthy breeding for our native orchids. In nectar-bearing plants, foraging insects will frequently move between flowers on the same plant and between neighbouring plants. Called “optimal foraging”, exhausting local nectar supplies in a patch before putting energy into finding a new buffet makes economic sense for a nectar-feeding insect. Sexual deception however, has been shown to drive pollinators far from the flower after being fooled, so that pollen escapes the local neighbourhood. As a plant, your neighbours are likely to be related to you, thus deception is a way of ensuring offspring quality by avoiding breeding with your relatives.

Another factor supporting the profusion of our sexually deceptive species is Australia’s immense diversity of insects to fool. Although there are examples of gnat and ant sexual deception systems, wasps are the most commonly targeted pollinator for our orchids. Incredibly, we are only now beginning to uncover the immense hidden diversity of Australian wasps. For example, a recent study in a small patch of bush near Margaret River uncovered 28 species of wasps, most of which were previously unknown to science. With each of these species most likely having their own private sex-pheromone cocktail, there is seemingly a kaleidoscope of chemical communication channels available for different orchids to exploit.

Despite our deepening understanding of the natural history of sexual deception, its repeated occurrence in Australia remains a true puzzle.

Rather than luring its pollinator with the promise of food this flower uses an equally, if not more, powerful motivator: sex. Undetectable to human senses, the orchid’s advertisement is a precise chemical mimicry of a female wasp’s sex pheromone. This is targeted marketing at its finest, as the use of a signature sex pheromone ensures that the orchid attracts only males of a specific species of wasp.

In 2010 I was fortunate to accompany herpetologist Dr. Renee Catullo on a trip through Australia’s Top End collecting little brown frogs. Here are some of my field notes along with photography. While my photography has vastly improved, I’m not sure I’ve felt so inspired to write since that trip. I hope this series of vignettes communicate some of the flavour and excitement of the wet season in Australia’s monsoon tropics.

I was relieving myself a stone’s throw from the inundated Carpentaria Highway when I saw my first Brolga. Its lanky form drifted indolently against a pastel sky as the tropical sun poured out late afternoon’s final slanting blaze. I finished my business, picked up my binoculars and shovel, tucked the bog roll under my arm, snatched a nearby burrowing frog with my free hand and wandered back to where Steve had left our hired Landcruiser: bogged up to the axles in the soft roadside sludge.

We had been following a map to the “Lost City”, scrawled on a take-away order pad by a gnarly-toothed cook at Cape Crawford’s “Heartbreak Hotel”. (This sentence is entirely true and gives me great pleasure to write). I had hoped to see the exceedingly elusive Carpentarian Grass-wren there, but like Grass-wrens, Lost Cities are difficult to find. Renee planned to make a 200 km push down and back the Tablelands Highway that night so our search for the Lost City was truncated in favour of beginning the evening’s driving. Our U-turn was truncated on the shoulder of the highway, which had been under floodwater for the last 3 or 4 days and needed little persuasion to engulf the hired Landcruiser.

We had been stalking 300 kms of the Carpentaria Hwy for the last two nights and had seen one car in that time. It was 5.30pm on Sunday and we had enough food and water in the car to last us a few nights should it come to that, but in a great fluke of variance the next 30 minutes sent us three different cars. Our eventual saviour rocked up in a big old Landcruiser. As a spry indigenous woman jumped out of the passenger seat to retrieve a rope from the tray, a man as weathered, wide and red as our fair country hopped out of the driver’s seat. Our saviour was in a hurry, the rope quickly linked our cars back to back and he barked some instructions to an unsure Steve behind the wheel of our stricken vehicle. Soon wheels were spinning as his truck revved and squealed, switching about like a hooked barramundi. Steve eventually found 4WD, low range, reverse and our steed was yanked backwards from the muck.

Impeded on the Carpentaria Hwy

That was the beginning of one of the most eventful evenings of the trip. We were now running a bit late, but counted ourselves lucky to have gotten away with an hour delay. The drive ahead saw us traveling south down the Tablelands Highway, a road that splits rocky cliffs of rust red upon which the ivory white bark of eucalypts is thrown into dramatic contrast by the light from the setting sun. We stopped on a vast flat grass plain after dark and watched the moon rise, a giant languid yellow disc that was to light the rest of our evening.

It is worth mentioning that I only refer to the roads by name because there are so damn few of them here in the Top End. Each road takes on a character through its interaction with the elements, the lansdscape, and the idiosyncrasies of the disparate roadhouses, towns and remote communities it links across vast distance. These bitumen ribbons of country and the hours spent on them forms a large part of the Top End experience for residents and visitors alike.

Soon after dark the animals started throwing themselves in front of the car. A remote wet season highway at night is chaos. I’ve never seen so much wildlife ever, and you’ll be pleased to know most of it survived. Bustards, nightjars, owls and curlews appear in the headlights, I brake and they make a flapping flash in my periphery as they fly within centimetres of the windscreen. Agile wallabies bound in groups across the road ahead, or sometimes they just stand on the road shoulder, waiting for us to drive closer before darting out into our path. The road is littered with frogs, only discernable as the car drives on over them. But one can’t look out for them as it is much more important to spot the groups of Brahman cattle from a distance safe enough to slow down. At several points we’re slowed to sub 10 km/h as dozens of cows slowly bumble down the road in front of you for hundreds of meters. There is nothing to do but travel along behind until they find a spot to vacate the bitumen. In addition to the fauna there’s water to watch for, the road is flooded in some points up close to a meter, and the floods have dragged onto the road a flotsam of logs, branches and general crap to drive around.

Tablelands Hwy denizen with biologist for scale – Black-headed python

When we swapped drivers I had run over what I was horrified to hear Renee later estimate as a total of 30 burrowing frogs. At the wheel, Steve the frog lover perhaps avoided a few more than I did, but unfortunately he was unable to avoid a nail tail wallaby. The nail tails are beautiful creatures up close, covered in soft, fine light caramel fur with faint white markings. A dark dorsal line runs from the back of their neck down their spine to the nail protruding from a spray of thick black bristles at the end of the tail.

In 2010 I was fortunate to accompany herpetologist Dr. Renee Catullo on a trip through Australia’s Top End collecting little brown frogs. Here are some of my field notes along with photography. While my photography has vastly improved, I’m not sure I’ve felt so inspired to write since that trip. I hope this series of vignettes communicate some of the flavour and excitement of the wet season in Australia’s monsoon tropics.

3 February 2010

Birds, wind, insects, all still, all silent. An atmosphere of exhaustion settles in as the temperature draws near 40. Wanda Inn roadhouse sits baking between red clay and the azure expanse of the outback sky. This would be home for the night, and we’d best get some sleep before tonight’s activities.

Littoria caerulea find respite from day’s dry heat by hanging out in the toilets.

The area hadn’t seen rain for a while but on dusk the moodily flashing cumulus clouds in the distance promised a change in the weather. Soon after dark we find ourselves in frogless country, standing on the roof of the Landcruiser captivated by the lightshow of distant electrical storms. As time passes the breeze blows harder, and we drive closer to its origin. Soon we are buffeted by a stiff cool wind, petrichor fills the air, fat drops of rain spatter and we are surrounded on all sides by the intermittent discharge of three kinds of lighting. Coruscating bolts crack somewhere between us and the horizon, leaving their jagged after-image in negative on our retinas, snaking fingers of electrical tracery race bifurcating and branching in the clouds overhead, and distant bolts diffused through cloud and rain suddenly bathe the land in white, turning puddles into mirrors for the silver sky before plunging us back into darkness. The weather is completely in character for the Northern Territory: at once utterly romantic and dangerously thrilling.

We stop periodically to stand on the car and take pictures of the gathering storm (still no frogs calling) but when a blinding bolt cracks too close for comfort we quickly curtail our roof-standing. Discouraged by the lack of frogs, the talk turns to gin and tonic at our next stop, but when we get there, rather than a refreshing drink, the distinctive chip of the Stonemason toadlet greets us and the frogging is on.

And so the wind blows, fat drops spatter our skin as we stride through waist high grass and negotiate ankle-twisting holes, boot-sucking puddles, and mud that cakes to the mud that’s caked on the mud caked on our boots. We wiggle between the slick ground and barbed wire fences while the sky flashes chaotically, rumbling and booming, and we stalk around pools thronging with the disorientating din of burrowing frogs, occasionally glimpsing the wide eyes of the other when the land in an instant becomes awash in the white of a nearby burst of lighting. This was trench-warfare frogging.